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Dissertation - HQ

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102 Vertical distribution during ontogeny<br />

Situate ontogeny<br />

among other factors<br />

Regression between<br />

size and depth<br />

Before exploring the differences between the distribution of ontogenetic<br />

stages, one must make sure that other sources of variability are<br />

not obscuring the potential effect of ontogeny. For example, post-flexion<br />

larvae may be always a few meters lower in the water column than<br />

pre-flexion larvae, but if both pre- and post-flexion larvae are within<br />

0-20 m when the thermocline is at 30 m and within 20-50 m when it<br />

is at 60 m, testing for a global difference in location through a rank<br />

test such as the Wilcoxon-Mann-Whitney test will show nothing. One<br />

solution is to work with the difference in z cm at each station, rather than<br />

with the z cm themselves. Another possibility, which gives additional<br />

information on the system, is to identify the other sources of variability<br />

and eliminate them before testing the effect of ontogeny.<br />

Successive regression trees were constructed to hierarchise the factors<br />

influencing the distribution of z cms, and situate ontogeny among<br />

those. The explanatory variables considered in addition to ontogeny<br />

were taxonomic (family), temporal (time of day), geographic (latitude,<br />

longitude, location with respect to the atoll, i.e. windward, leeward),<br />

and hydrographic (depth of thermo-, halo-, pycnoclines, and of the<br />

fluorometry maximum, mean current speed in the surface layer). When<br />

several factors were correlated (e.g. depth of thermo- and pycnocline)<br />

they were tested independently and only the most explanatory was<br />

kept in the final tree. For discrete explanatory variables, the effect of<br />

influential factors was investigated by comparing z cms between groups<br />

(by taxon, by ontogenetic stage, etc.) using non-parametric tests for<br />

differences in medians (Wilcoxon-Mann-Whitney and Kruskal-Wallis).<br />

Homogeneity in variances was tested using the Fligner-Killeen test.<br />

When variances were different between groups, the choice was made<br />

to still use the same tests but to lower the significance level to 0.01, to<br />

account for the higher risk of α-error (distributions were clearly nonsymmetric,<br />

preventing the use of robust rank procedures, as mentioned<br />

in the section “The problem of unequal variances”, page 99). The effect<br />

of continuous explanatory variables was estimated by regression using<br />

Generalised Linear Models with a gamma distribution of errors.<br />

Over 3000 larvae were measured and their size was used as a proxy<br />

for development. Indeed larvae usually reach a particular ontogenetic<br />

level at a given size rather than at a given age 64 . They allowed to test<br />

whether there was a continuous change in vertical distribution during<br />

ontogeny (i.e. along with increasing sizes) through a regression analysis.<br />

Of course size varies greatly among different fish taxa. Therefore, sizes<br />

were normalised per taxon, i.e. for each of the lowest taxonomic units<br />

identified, the size of the smallest fish captured was set to zero while the<br />

size of the largest fish was scaled to one. While the ranges of ontogenetic<br />

stages captured probably differed between taxa (i.e. size = 1 did not<br />

correspond to the same point in development for all taxa), this brought<br />

sizes on a more homogenous scale. Relative size and depth of capture<br />

could then be compared. However, because of the patchiness in the<br />

distribution of larvae, two larvae of the same taxon captured at the

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